1 08-F Sentidos Somaticos - PDF 2

Machine Translated by Google
Sensory Systems
Sensory Systems
OpenStax Anatomy & Physiology 2nd ed

Sensory Systems
Range from simple neurons to complex sense organs
Sensory receptors transduce incoming stimuli into changes in membrane potential (local graded potentials)
Sensory Systems
Sensory Systems
Sensitivity to Multiple Modalities
• Adequate stimulus – preferred or most sensitive stimulus

modality (many receptors can also be excited by other
stimuli, if sufficiently large, eg, pressure on eyelid
perceive bright light)
• Polymodal receptors – naturally sensitive to more than one

stimulus modality, eg, ampullae of Lorenzini in sharks
(electroreception, magnetoreception, thermoreception)
• Nociceptors – sensitive to strong stimuli, eg, pain; many are

polymodal receptors
Sensory Systems
Classification:
Based on stimulus location:
• Telereceptors – distant stimuli (vision, hearing) •
Exteroceptors – stimuli on the outside of the
body, (pressure, temperature)
• Interoceptors – stimuli inside the body (blood
pressure, blood oxygen)
Based on stimulus type:

• Chemoreceptors •
Mechanoreceptors •
Photoreceptors •
Thermoreceptors •
Osmoreceptors •
Electroreceptors •
Magnetoreceptors
Sensory Systems
Graded Potentials Sensory Systems

•Generator potential – sensory receptor is also the primary afferent neuron

•Receptor potential – sensory receptor is separate from the afferent neuron
GRADED POTENTIAL
POTENTIAL ACTION GRADED POTENTIAL
POTENTIAL ACTION
Sensory Systems
GRADED POTENTIALS
• Short-lived, localized changes in membrane potential
• Depolarizations (EPSPs) or hyperpolarizations (IPSPs)

• Graded potential spreads as local currents change the membrane potential of adjacent regions (electrotonic conduction)
• Occur when a stimulus causes gated ion channels to open
• Eg, receptor potentials, generator potentials, postsynaptic potentials
• Magnitude varies directly (graded) with stimulus strength
• Decrease in magnitude with distance as ions flow and diffuse through leakage channels
• Short-distance signals
Membrane
potential
(mV)
Stimulus
Depolarized region
–70
Plasma
membrane
Distance (a few mm)
(a) Depolarization: A small patch of the (b) Spread of depolarization: The local currents (black (c) Decay of membrane potential with distance: Because current is lost
membrane (red area) has become depolarized. arrows) that are created depolarize adjacent through the leaky plasma membrane, the voltage declines with
membrane areas and allow the wave of distance from the stimulus (the voltage is decremental).
depolarization to spread. Consequently, graded potentials are short-distance signals.
Sensory Systems
• A single EPSP cannot induce an action potential
• EPSPs can summate to reach threshold
• IPSPs can also summate with EPSPs, canceling each other out
Temporary summation
One or more presynaptic neurons transmit impulses in rapid-fire order
Spatial summation
Postsynaptic neuron is stimulated by a large number of terminals at the same time
E1 E1 E1 E1
E2 I1
Threshold of axon of
postsynaptic neuron
Resting potential
E1 E1 E1 E1 E1 + E2 I1 E1 + I1
time time time time
(a) Non-summation: (b) Temporary summation: (c) Spatial summation: (d) Spatial summation of
2 simultaneous stimuli at EPSPs and IPSPs:
2 stimuli separated in time cause 2 excitatory stimuli close
EPSPs that do not in time cause EPSPs
different locations cause Changes in membane
add together. that add together.

EPSPs that add together. potential can cancel each other
out.
Excitatory synapse 1 (E1)

Inhibitory synapse (I1)
Sensory Systems
(1) Somatosensory system: (2) Special senses:
• (a) Exteroceptive (mechanoceptive, thermoceptive and nociceptive) • Vision
(b) Proprioceptive (positional) • Audition / Equilibrium
Olfaction
(c) Enteroceptive (viscerosensory) Taste
(to) (c)
(b)
Receptor Location Somatosensory System - Exteroceptive
The somatosensory system provides

information to the central nervous
system (CNS) about the state of the
body and its contact with the world.
Integrated by sensory receptors that

transduce mechanical (pressure,
stretch, and vibrations) and thermal
energies into electrical signals.
These electrical signals are called

generator or receiver potentials and
occur in the distal ends of axons of first-order
somatosensory neurons, where
they trigger action potential trains that
reflect information about the
characteristics of the stimulus.
The cell bodies of these neurons are

located in dorsal root and cranial nerve
ganglia.
Sensory organ Primary stimuli Location Structure adaptation

Until recently, ASIC (acid-sensing ion channel) proteins,

which belong to the ENaC/DEG superfamily, had been
thought to be the channel proteins underlying the
cutaneous mechanoactivated currents.

Currently, Piezo2 is thought to be channel protein

underlying the transduction for cutaneous mechanical
rapidly adapting responses
It forms a nonselective cation pore that opens in response

to mechanical stimuli.

Currently, Piezo2 is thought to be channel protein

underlying the transduction for cutaneous mechanical
rapidly adapting responses
It forms a nonselective cation pore that opens in response

to mechanical stimuli.

Biophysical Reviews (2022) 14:15–20
https://doi.org/10.1007/s12551-022-00935-9
COMMENTARY
2021 Nobel Prize for mechanosensory transduction

Boris Martinac1
Received: 15 December 2021 / Accepted: 2 February 2022 / Published online: 19 February 2022
© The Author(s) 2022
Abstract
The shared Nobel Prize for Physiology or Medicine 2021 Written by someone who has worked in the mechanobiology field for close to 40 years, this commentary describes some historical background to the
recent award of one-half of the Nobel Prize for Physiology or Medicine to Ardem Patapoutian for his discovery of the family of mechanosensitive Piezo
awarded to David Julius and Ardem Patapoutian for ion channels, which function as mechanoreceptors sensing the environment in senses such as touch, pain, and proprioception.
their respective breakthrough discovery of the vertebrate Keywords Mechanosensitive ion channels · MscL · MscS · Piezo1 · Piezo2
thermosensory TRP and mechanosensory Piezo ion The shared Nobel Prize for Physiology or Medicine 2021 awarded to Brief history
David Julius and Ardem Patapoutian for their respective breakthrough
channels recognizes the significance of the evolutionarily discovery of the vertebrate thermo-sensory TRP and mechanosensory
Piezo ion channels recog-nizes the significance of the evolutionarily
The existence of mechanosensitive ion channels was first
postulated in 1950 by Bernard Katz, who won the Nobel Prize for
inherent ability that living organisms, from bacteria to humans, possess in Physiology or Medicine in 1970 for his work on nerve physiology (Katz
inherent ability that living organisms, from bacteria to sensing and responding to changes in their surrounding environment. 1950). A quarter of a century later, Georg von Békésy, the 1961 Nobel
This commentary is restricted to the prize relating to mechanosensitive Prize winner for Physiology or Medicine, discussed a possible existence
humans, possessing in sensing and responding to changes ion channels. of a mechanical receptor in frequency discrimination in the ear (Von
Bekesy 1974). Several years later, Corey and Hudspeth suggested the
Given the essential role of water for the existence of life and the existence of vertebrate mechanosensitive ion channels in bullfrog cochlear
in their surrounding environment. presence of osmotic forces throughout the evolution of different life forms hair cells (Corey and Hudspeth 1979). Shortly after, thanks to the advent
on Earth, mechanosensitive ion channels may be the oldest type of of the patch clamp technique (Hamill et al. 1981), the first single mecha-
mechanoreceptors that evolved as primary signaling molecules supporting nosensitive ion channels were recorded from chick skeletal muscle by
mecha-nosensory physiology of living organisms. Without the ability to Guharay and Sachs (Guharay and Sachs 1984), from innervated muscle
perceive sensations of touch, hearing, sight, taste, smell, temperature, or of Xenopus laevis by Brehm and collages (Brehm et al. 1984) and from
pain, the outside world would cease to exist for vertebrate organisms, giant spheroplasts of E. coli by Martinac and colleagues (Martinac et al.
including humans, which emphasizes the importance of sensory input for 1987).
the existence of life . To this point, Piezo2 members of the Piezo ion
channel family serve as mechanoreceptors sensing the environment in
senses such as touch, pain, and proprioception (Ernfors et al. 2021) (Fig. It is to the credit of the Nobel Assembly, who duly recognized the
1). pioneering studies on bacterial mechanosensitive channels (Fig. 2), which
firmly established the existence of mechanosensitive channels in the late
1980s (Cox et al.
2018; Delcour et al. 1989; Martinac et al. 1987; Sukharev et al. 1994).
This collective work ultimately paved the way to this year's Nobel Prize
for the discovery of the Piezo mecha-noreceptor family (Ernfors et al.
2021).
* Boris Martinac
b.martinac@victorchang.edu.au
1
Molecular Cardiology and Biophysics Division, Victor Chang
Cardiac Research Institute and St. Vincent's Medical School, Darlinghurst,
NSW 2010, Australia
Vol.:(0123456789) 1 3
Somatosensory System - Proprioceptive



Somatosensory System - Nociceptive
The axons that carry painful and thermal sensations are members of the relatively slowly conducting Aÿ
and C classes, and originate mostly as “free nerve endings.”
Aÿ axons conduct signals faster (fast pain: sharp, acute) than C fibers (slow pain: dull, diffuse).
Afferent fibers are classified according to their sensitivity to mechanical, thermal, and chemical stimuli:
• C mechanosensitive
• C mechanoheat-sensitive (polymodal)
• C cold-sensitive
• Aÿ cold-sensitive
• Aÿ mechanosensitive
• Aÿ mechanoheat-sensitive
Receptor Location Somatosensory System - Nociceptive

Somatosensory System - Thermoceptive

Receiver Location
Biophysical Reviews (2022) 14:15–20
https://doi.org/10.1007/s12551-022-00935-9
COMMENTARY
2021 Nobel Prize for mechanosensory transduction

Boris Martinac1
Received: 15 December 2021 / Accepted: 2 February 2022 / Published online: 19 February 2022
© The Author(s) 2022
Abstract
The shared Nobel Prize for Physiology or Medicine 2021 Written by someone who has worked in the mechanobiology field for close to 40 years, this commentary describes some historical background to the
recent award of one-half of the Nobel Prize for Physiology or Medicine to Ardem Patapoutian for his discovery of the family of mechanosensitive Piezo
awarded to David Julius and Ardem Patapoutian for their ion channels, which function as mechanoreceptors sensing the environment in senses such as touch, pain, and proprioception.
respective breakthrough discovery of the vertebrate Keywords Mechanosensitive ion channels · MscL · MscS · Piezo1 · Piezo2
thermosensory TRP and mechanosensory Piezo ion The shared Nobel Prize for Physiology or Medicine 2021 awarded to Brief history
David Julius and Ardem Patapoutian for their respective breakthrough
channels recognizes the significance of the evolutionarily discovery of the vertebrate thermo-sensory TRP and mechanosensory
Piezo ion channels recog-nizes the significance of the evolutionarily
The existence of mechanosensitive ion channels was first
postulated in 1950 by Bernard Katz, who won the Nobel Prize for
inherent ability that living organisms, from bacteria to humans, possess in Physiology or Medicine in 1970 for his work on nerve physiology (Katz
inherent ability that living organisms, from bacteria to sensing and responding to changes in their surrounding environment. 1950). A quarter of a century later, Georg von Békésy, the 1961 Nobel
This commentary is restricted to the prize relating to mechanosensitive Prize winner for Physiology or Medicine, discussed a possible existence
humans, possessing in sensing and responding to changes ion channels. of a mechanical receptor in frequency discrimination in the ear (Von
Bekesy 1974). Several years later, Corey and Hudspeth suggested the
Given the essential role of water for the existence of life and the existence of vertebrate mechanosensitive ion channels in bullfrog cochlear
in their surrounding environment. presence of osmotic forces throughout the evolution of different life forms hair cells (Corey and Hudspeth 1979). Shortly after, thanks to the advent
on Earth, mechanosensitive ion channels may be the oldest type of of the patch clamp technique (Hamill et al. 1981), the first single mecha-
mechanoreceptors that evolved as primary signaling molecules supporting nosensitive ion channels were recorded from chick skeletal muscle by
mecha-nosensory physiology of living organisms. Without the ability to Guharay and Sachs (Guharay and Sachs 1984), from innervated muscle
perceive sensations of touch, hearing, sight, taste, smell, temperature, or of Xenopus laevis by Brehm and collages (Brehm et al. 1984) and from
pain, the outside world would cease to exist for vertebrate organisms, giant spheroplasts of E. coli by Martinac and colleagues (Martinac et al.
including humans, which emphasizes the importance of sensory input for 1987).
the existence of life . To this point, Piezo2 members of the Piezo ion
channel family serve as mechanoreceptors sensing the environment in
senses such as touch, pain, and proprioception (Ernfors et al. 2021) (Fig. It is to the credit of the Nobel Assembly, who duly recognized the
1). pioneering studies on bacterial mechanosensitive channels (Fig. 2), which
firmly established the existence of mechanosensitive channels in the late
1980s (Cox et al.
2018; Delcour et al. 1989; Martinac et al. 1987; Sukharev et al. 1994).
This collective work ultimately paved the way to this year's Nobel Prize
for the discovery of the Piezo mecha-noreceptor family (Ernfors et al.
2021).
* Boris Martinac
b.martinac@victorchang.edu.au
1
Molecular Cardiology and Biophysics Division, Victor Chang
Cardiac Research Institute and St. Vincent's Medical School, Darlinghurst,
NSW 2010, Australia
Vol.:(0123456789) 1 3
Thermoceptors belong to the TRP (transient receptor potential) family of ionic channels

Receptor Location Somatosensory System - Pathways
Dorsal column-ML tract

(epicritic pathway)
Spinothalamic tract
(protophatic pathway)
Somatosensory System - Processing

Receiver Location
Signal processing starts at the receiver level
Sensory receivers must encode four types of information • Stimulus

modality • Stimulus location
• Stimulus intensity •
Stimulus duration
Contrast is enhanced at the receiver field

• Lateral inhibition

Receiver Location
Modality
Theory of labeled lines: Specific neuroanatomic pathways from the sensory cells to the integrating
centers (polymodal receptors are exceptions: encode modality via temporal patterns of APs)
Location
Region of the sensory surface that generates a response when stimulated
Smaller receptive field à more precise location à greater acuity
Intensity
Receptor potential amplitude is proportional to stimulus intensity
Action potential frequency is proportional to stimulus intensity
Summation: •
Spatial: using more than one sensory receptor cell • Temporal:
concurrent inputs increase signal strength
Duration •
Tonic receptors: Slow adapting • Phasic
receptors: Fast adapting
Lateral inhibition
Signals from neurons at the center of the stimulated area inhibit neurons on the periphery

Receiver Location
Modality
Theory of labeled lines: Specific neuroanatomic pathways from the sensory cells to the integrating centers

Receiver Location
Location

Receiver Location
Location

Receiver Location
Location

Receiver Location
Location
Modality
Theory
Region of
of labeled
the sensory
lines:surface
Specificthat
neuroanatomic
generates a response
pathwayswhen
from the
stimulated
sensory cells to the integrating centers

Receiver Location
Intensity
Receptor potential amplitude is proportional to stimulus intensity
Action potential frequency is proportional to stimulus intensity

Receiver Location
E1 E1 E1 E1
E2 I1
Threshold of axon of
postsynaptic neuron
Resting potential
E1 E1 E1 E1 E1 + E2 I1 E1 + I1
time time time time
(a) Non-summation: (b) Temporary summation: (c) Spatial summation: (d) Spatial summation of
2 simultaneous stimuli at EPSPs and IPSPs:
2 stimuli separated in time cause 2 excitatory stimuli close
EPSPs that do not in time cause EPSPs different locations cause Changes in membane
add together. that add together. EPSPs that add together. potential can cancel each other
out.

• A single EPSP cannot induce an action potential
• EPSPs can summate to reach threshold
Inhibitory synapse (I1)
• IPSPs can also summate with EPSPs, canceling each other out
Temporary summation
One or more presynaptic neurons transmit impulses in rapid-fire order
Spatial summation
Postsynaptic neuron is stimulated by a large number of terminals at the same time
Intensity
Summation:
• Spatial: using more than one sensory receptor cell
• Temporary: concurrent inputs increase signal strength

Receiver Location
Intensity
Summation: •
Spatial: using more than one sensory receptor cell • Temporal:
concurrent inputs increase signal strength
Somatosensory
Tonic and PhasicSystem - Processing
Receptors, Cont.
Two classes of receptors that encode stimulus duration

• Phasic – produces APs only at the beginning or end of the stimulus •
Tonic – produces APs as long as the stimulus continues
Duration
• Tonic receptors: Slow adapting •
Phasic receptors: Fast adapting

Receiver Location
Lateral inhibition
Signals from neurons at the center of the stimulated area inhibit neurons on the periphery
Receiver Location
Receiver Location
Receptor Location Somatosensory System
Action potentials code stimulus intensity through changes in

frequency, eg, strong stimuli à high frequency
• Dynamic range – range of intensities for which receivers

can encode stimuli
• Threshold detection – weakest stimulus that produces a
response in a receiver 50% of the time
• Saturation – top of the dynamic range; all available
proteins have been stimulated
Relationships between stimulus intensity and AP frequency
• (A) Linear across large range of intensities: large change in

stimulus causes a small change in AP frequency à
large dynamic range, poor sensory discrimination
• (C) Linear across small range of intensities: small change in
stimulus causes a large change in AP frequency à
small dynamic range, high sensory discrimination
• (B) Encode a wide range of stimulus intensities using a
single receptor cell (logarithmic response)
Fine discrimination at certain intensities (eg, low)
Coarse discrimination at other intensities (eg, high)
Range fractionation – groups of receptors work together to increase dynamic

range without decreasing sensory discrimination
Transform mechanical stimuli into electrical signals

All organisms and cells can sense and respond to mechanical stimuli
Two main types
• ENaC – epithelial sodium channels •
TRP – transient receptor potential
Touch and Pressure

Three classes
• Baroreceptors – interoceptors that detect pressure changes
• Tactile receptors – exteroceptors that detect touch, pressure, and vibration on the body surface
• Proprioceptors – monitor the position of the body
Insects
Two types of mechanoreceptors
Table 7.1
Type 1 – External Surface
Two common types of

sensilla
• Trichoid – hairlike
• Campaniform –
bell-shaped
Figure 7.13
Type 1 – Internal Surface

• Scolopidia – bipolar neuron
and complex accessory
cell (scolopale)
• Can be isolated or
grouped to form
chordotonal organs
• Most function in
proprioception
• Can be modified into
tympanal organs for sound
detection
Figure 7.14
Vertebrate Tactile Receptors

• Widely dispersed
• Function as isolated sensory cells
• Free nerves ending or enclosed in
accessory structures (eg, Pacinian
corpuscle)
Figure 7.15
Proprioceptors
Monitor the position of the body
Three major groups
• Muscle spindles – located on the surface of the muscle
and monitor muscle length
• Golgi tendon organs – located at the junction between
skeletal muscles and tendons and monitor tendon tension
• Joint capsule receptors – located in the capsules

that enclose joints and detect pressure, tension,
and movement in the joint
Thermoreception
Central thermoreceptors – located in the

hypothalamus and monitor internal
temperature
Peripheral thermoreceptors – monitor
environmental temperature
• Warm-sensitive
• Cold-sensitive
• Thermal nociceptors – detect painfully hot
stimuli
ThermoTRPs – TRP ion channel
thermoreceptor proteins
Specialized Thermoreception
Specialized organs for detecting heat
radiating objects at a distance
Pit organs – pit found between the eye and
the nostril of pit vipers
Can detect 0.003°C changes (0.5°C for
humans)
Magnetoreception
• Ability to detect magnetic fields
• eg, migratory birds, homing salmon
• Neurons in the olfactory epithelium of
rainbow trout contain particles that
resemble magnetite

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Machine Translated by Google

OpenStax Anatomy & Physiology 2nd ed

• Adequate stimulus – preferred or most sensitive stimulus

• Polymodal receptors – naturally sensitive to more than one

• Nociceptors – sensitive to strong stimuli, eg, pain; many are

Based on stimulus type:

Graded Potentials Sensory Systems

Range from simple neurons to complex sense organs

•Generator potential – sensory receptor is also the primary afferent neuron

POTENTIAL ACTION GRADED POTENTIAL

• Short-lived, localized changes in membrane potential

• Depolarizations (EPSPs) or hyperpolarizations (IPSPs)

time time time time

add together. that add together.

Excitatory synapse 1 (E1)

Receptor Location Somatosensory System - Exteroceptive

The somatosensory system provides

Integrated by sensory receptors that

These electrical signals are called

The cell bodies of these neurons are

Receptor Location Somatosensory System - Exteroceptive

Sensory organ Primary stimuli Location Structure adaptation

Receptor Location Somatosensory System - Exteroceptive

Until recently, ASIC (acid-sensing ion channel) proteins,

Receptor Location Somatosensory System - Exteroceptive

Until recently, ASIC (acid-sensing ion channel) proteins,

Currently, Piezo2 is thought to be channel protein

It forms a nonselective cation pore that opens in response

Receptor Location Somatosensory System - Exteroceptive

Until recently, ASIC (acid-sensing ion channel) proteins,

Currently, Piezo2 is thought to be channel protein

It forms a nonselective cation pore that opens in response

Receptor Location Somatosensory System - Exteroceptive

2021 Nobel Prize for mechanosensory transduction

Somatosensory System - Proprioceptive

Somatosensory System - Proprioceptive

Somatosensory System - Proprioceptive

Somatosensory System - Nociceptive

Receptor Location Somatosensory System - Nociceptive

Somatosensory System - Thermoceptive

2021 Nobel Prize for mechanosensory transduction

Somatosensory System - Thermoceptive

Somatosensory System - Thermoceptive

Receptor Location Somatosensory System - Pathways

Dorsal column-ML tract

Somatosensory System - Processing

Signal processing starts at the receiver level

Sensory receivers must encode four types of information • Stimulus

Contrast is enhanced at the receiver field

Somatosensory System - Processing

Somatosensory System - Processing

Somatosensory System - Processing

Somatosensory System - Processing

Somatosensory System - Processing

Somatosensory System - Processing

Somatosensory System - Processing

Somatosensory System - Processing

time time time time

Excitatory synapse 1 (E1)

Somatosensory System - Processing

Two classes of receptors that encode stimulus duration

Somatosensory System - Processing